Colony fission affects kinship in a social insect

Establishment of new groups is an important step in the life history of a social species. Fissioning is a common mode not only in group proliferation, for instance, as a regular part of the life cycle in the honey bee, but also when multiple females reproduce in the same group, as in multiple-queen ant societies. We studied the genetic consequences of fissioning in the ant Proformica longiseta, based on DNA microsatellites. In P. longiseta, new nests arise by fissioning from the old ones when they grow large, and the daughter nests consist of workers and queens or queen pupae but never both. Our results show that fissioning is not entirely random with respect to kinship. Workers tend to segregate along kin lines, but only when the initial relatedness in the parental nests is low. Workers in a daughter nest also tend to be associated with closely related adult queens, whereas such an association is not detected between workers and queen pupae. Most queens and workers are carried to the daughter nest by a specialized group of transporting workers, suggesting active kin discrimination by them. Fissioning pattern in P. longiseta is different from that found in other social insects with regular fission (e.g., the honey bee, swarm-founding wasps), where no fissioning along kin lines has been found. It does, however, resemble fissioning in another group of social animals: primates.     

Influence of social factors on sex ratio at birth,maternal investment and young survival in a prosimian primate

Summary In order to determine whether social factors influence sex ratio at birth in lesser mouse lemurs, experiments were conducted during 5 successive breeding periods on 51 females. At the beginning of the breeding season, females were either isolated (I) or grouped (G) in heterosexual groups with an increasing number of females (2, 3 or 4). To ensure mating, I females were introduced in a group only during the oestrous period. After mating, both I and G females were isolated during pregnancy and lactation. Reproductive capacities of females in terms of oestrus occurrences (n = 324), impregnations (n–210), pregnancies (n = 136) or abortions (n = 38) or litter sizes (1–3 young) were affected neither by age and parity of females nor by group housing prior to conception. G females produced significantly more sons than daughters (67% males for 189 newborn) while females living alone except during the mating period demonstrated a significant inverse tendency (39.6% males for 96 newborn). Distribution of sexes in litters was statistically different from random and varied according to the shift of sex ratio at birth. In G females, the shift in the sex ratio towards males was consistent across the different groups, independent of the number of females living together, suggesting that the presence of only 1 female is sufficient to induce a bias in the sex ratio. No correlation was found between infant survival at weaning and age, parity or group housing of the mother. The maternal investment allocated to male or female newborn was similar provided the litter contained at least 1 male. In litters without males, growth and survival of female infants were significantly less. These results on sex ratio bias in captive female mouse lemurs agree with directions of bias predicted by the local resource competition model for facultative sex ratio adjustment (Clark 1978). Nevertheless, the pattern observed in mouse lemurs appears to be independent of the nutritional state of the female and of differential maternal investment.     

Infant parking and nesting, not allomaternal care, influence Malagasy primate life histories

Allomaternal care is a rare, though phylogenetically widespread, mammalian infant care strategy. Among primates, the effects of allomaternal care are marked; its presence correlates with faster infant growth, younger age at weaning, and shorter interbirth intervals. Recent comparative research has found that such fertility benefits are absent in other mammals and are thus unique to primates. In large part because data describing lemur allomaternal care were lacking, the reproductive advantages of allomaternal care have never been demonstrated in Malagasy strepsirrhines. Using newly available data and rigorous phylogenetic methods, we extend this hypothesis to strepsirrhines and test whether allomaternal care in lemurs confers similar maternal reproductive benefits. Contrary to expectations, the presence of allomaternal care did not significantly impact lemur reproductive output; we did not find relationships between allomaternal care and either fetal or postnatal growth rates or interbirth intervals. Rather, infant parking and nesting, strategies employed primarily by litter-bearing species, were positively associated with faster fetal and postnatal infant growth, while nesting was negatively associated with interbirth interval. Thus, although each form of haplorrhine allomaternal care is also observed in Malagasy primates, the effects that these behaviors have on female reproductive output more closely resemble nonprimate mammals. We suggest that Malagasy strepsirrhines may not equally benefit from allomaternal care compared to haplorrhines because reproductive rates are less flexible and allomaternal care may instead increase infant survival in Madagascar’s harsh and unpredictable environment. Our study has significant implications for understanding the evolution of infant care and developmental trajectories in mammals.     

Solitary nesting and reproductive success in the paper wasp Polistes aurifer

Female Polistes paper wasps are capable of independent nesting, yet many populations demonstrate a mixture of solitary and cooperative nest foundation. Previous studies of Polistes have found survival and/or productivity advantages of cooperative nest foundation compared to solitary nesting, and reproductive skew models have been designed to predict the dynamics of such flexible cooperation. In this paper, we examine the success of different nesting strategies in a previously unstudied population of Polistes aurifer in southern California. The colony cycle of this population is less synchronous than that of other temperate species, and the frequency of solitary nesting averages 86.2%. Our data suggest that this low rate of cooperative nest founding is adaptive, as demonstrated by the lack of survival or productivity advantages for cooperative foundress associations. Due to foundress turnover and nest foundation later in the season, many nests produce only one set of offspring. This results in a loss of the eusocial nature of some nests in the population. Data from a small sample of multifoundress nests show significant positive reproductive skew, despite concession model predictions that skew should be low in populations with low ecological constraints on independent nesting. This lack of support for the concessions skew model reflects a diminished incentive for cooperation.Communicated by L. Keller     

Colonial and single breeding in fieldfares,Turdus pilaris L.: a comparison of nesting success in early and late broods

Summary Pairs of fieldfares breed singly or in colonies. Their breeding season was subdivided into an early part, when the trees were still leafless, and a late part, when the trees had leaves. Early colonial pairs has a significantly higher nesting success than single pairs. However, amongst late broods the nesting success of both categories of pairs was the same. Colonies of only two pairs show a weak trend suggesting that it could be advantageous for early broods to have a neighbour nearby, while the nesting success of late broods seems to be highest if the neighbour is farther away. The number of neighbours is more important than the distance to the nearest neighbour when colonies of different sizes are examined. The nesting success of colonial broods is also influenced by the time of breeding. Success increases with colony size, in early colonies, but the opposite applies for the late breeding season.A little owl was placed near single and colonial nests to show how communal defence contributed to the greater success of early colonial nests. Predators entering a colony risk being contaminated by significantly more faeces due to the higher rate of attacks by fieldfares using aimed defecation near colonial nests and these could affect the predators' flying ability.The possibility of fieldfare colonies serving as information centres for food finding is also discussed.     

Philopatry,reproductive success of females,and maternal investment in the red-necked wallaby

Summary Female red-necked wallabies settle within their mothers' home ranges, apparently for life, while males disperse at about two years of age. However, sons spend much more time with their mothers before dispersing than do daughters of similar ages. Females who associate regularly with their subadult offspring are less likely to reproduce successfully at their next breeding attempt than are females who spend little time with their subadults, and sons therefore impose greater short-term reproductive costs on their mothers than do daughters. Females who are generally gregarious also suffer reduced reproductive success, even though reproductive success is independent of local density. It is suggested that the reproductive costs to females of associating with their subadult offspring, and other relatives, are incurred through tolerance of ecological competition from those kin, and therefore reflect a form of prolonged maternal investment, which is initially heaviest in sons but is sustained for longer periods in daughters. Females produce equal numbers of male and female offspring, and spend equal amounts of time suckling them in infancy.     

Changes in maternal investment with climate moderate social behaviour in a facultatively social bee

Behavioral Ecology and Sociobiology - Scents play an important role in the life of most terrestrial mammals and may transmit valuable information about conspecifics. Olfaction was long considered...     

Effects of temperature and precipitation on grassland bird nesting success as mediated by patch size

Grassland birds are declining faster than any other bird guild across North America. Shrinking ranges and population declines are attributed to widespread habitat loss and increasingly fragmented landscapes of agriculture and other land uses that are misaligned with grassland bird conservation. Concurrent with habitat loss and degradation, temperate grasslands have been disproportionally affected by climate change relative to most other terrestrial biomes. Distributions of grassland birds often correlate with gradients in climate, but few researchers have explored the consequences of weather on the demography of grassland birds inhabiting a range of grassland fragments. To do so, we modeled the effects of temperature and precipitation on nesting success rates of 12 grassland bird species inhabiting a range of grassland patches across North America (21,000 nests from 81 individual studies). Higher amounts of precipitation in the preceding year were associated with higher nesting success, but wetter conditions during the active breeding season reduced nesting success. Extremely cold or hot conditions during the early breeding season were associated with lower rates of nesting success. The direct and indirect influence of temperature and precipitation on nesting success was moderated by grassland patch size. The positive effects of precipitation in the preceding year on nesting success were strongest in relatively small grassland patches and had little effect in large patches. Conversely, warm temperatures reduced nesting success in small grassland patches but increased nesting success in large patches. Mechanisms underlying these differences may be patch‐size‐induced variation in microclimates and predator activity. Although the exact cause is unclear, large grassland patches, the most common metric of grassland conservation, appears to moderate the effects of weather on grassland‐bird demography and could be an effective component of climate‐change adaptation.     

Biparental investment and reproductive success in a subsocial desert beetle: the role of maternal effort

Parastizopus armaticeps is a nocturnal subsocial detritivorous desert tenebrionid that produces very few offspring per brood. The two environmental factors that constrain reproduction, rapid sand desiccation rate and food scarcity, are countered by biparental effort. Males dig and extend breeding burrows, maintaining their moisture level; females forage on the surface at night for high-quality detritus, the larval food. This was shown to be a scarce and unpredictable resource for which there is high competition. When food was supplemented in a field experiment, offspring number and survivorship doubled and burrow failure due to desiccation dropped from approximately half, the typical failure rate for unsupplemented burrows, to zero. Food supplementation did not, however, increase larval foodstore size and there was no difference in the size of the offspring produced. Supplemented females reallocated their time, foraging less and digging more with the male. This change in maternal behaviour patterns resulted in deeper burrows which remained moist longer, thus extending the larval production period. Female foraging efficiency, particularly food retrieval speed, determined how much time females could allocate to digging, consequently increasing the reproductive success of the pair. Burrow depth and sand moisture level at the burrow base were the major correlates of reproductive success, but the scarcity and unpredictability of high-quality food on the surface and the competition for this resource influenced the number of offspring indirectly through their effect on female behaviour. Received: 29 November 1996 / Accepted after revision: 7 December 1997     

Communal nursing in the evening bat,Nycticeius humeralis

Summary Nursing observations over two summers involving 76 lactating female evening bats, Nycticeius humeralis, and 128 pups in an attic in northern Missouri indicate that communal nursing occurs rarely until 2 weeks before weaning during which time over 18% of nursing bouts involve nondescendant offspring. The average relatedness among female pairs nursing non-descendant offspring, based on identity-by-descent estimates using allozyme data, was 0.04 (SE=0.12). Mitochondrial DNA d-loop sequence comparisons confirm that at most only 2 of 20 female pairs nursing non-descendant offspring came from the same matriline. Thus, females do not nurse matrilineal kin preferentially despite female natal philopatry. In addition, the average degree of relatedness within a colony (r=0.01, SE =0.03) is too low to provide any indirect benefits from communal nursing. Female error alone is insufficient to explain these observations because females tended to allow female nondescendant young to nurse but excluded nondescendant males, particularly when they had all-male litters. Furthermore, communal nursing bouts did not differ in duration from parental nursing bouts and involved 31 % of all banded females and 24% of all banded pups observed nursing. Communal nursing occurred most frequently when pups began hunting on their own and when lactating females attained their lowest average pre-fed body weight. Mortality during this period was higher for male than female pups, and relative weights implicate starvation as the cause. Time-lapse video records of four families of bats in captivity showed that the number of nursing bouts was proportional to daily weight change. I propose that these results are consistent with both immediate and delayed benefits accruing to females which experience variable hunting success. If a female with extra milk reduced her weight by dumping milk prior to her next foraging trip, she could obtain an immediate energetic benefit and maintain maximum milk production. By restricting such milk donations to nondescendant females she may also increase colony size and thereby enhance her future acquisition of information about foraging and roosting sites.     

Filial cannibalism and reproductive success in the maternal mouthbrooding cichlid fish Pseudocrenilabrus multicolor

Summary Pseudocrenilabrus multicolor is an iteroparous maternal mouthbrooding cichlid fish living in small and shallow waters in East Africa. It is subjected to large seasonal variations of population density. Laboratory experiments show that with increasing density and time after spawning the number of offspring present in the mouth decreased. By experimentally varying density and the numbers of fertilized and unfertilized eggs in the mouth at the end of spawning the following causes of brood size reduction were identified: (1) At high densities some of the spawned eggs are stolen by conspecifics before they are taken into the mouth. Of the eggs taken up a considerable proportion is unfertilized. (2) Unfertilized eggs are selectively swallowed from day 2 to day 5 after spawning. (3) Some of the fertilized eggs are also swallowed during mouthbrooding. (4) If the number of eggs in the mouth falls below a critical value of approximately 20% of the number of eggs spawned, they are all swallowed. By contrast, predation by conspecifics is insignificant as long as the brood is in the mother's mouth. The causes and functions of the mother's filial cannibalism are discussed.     

Prenatal social conditions shape offspring adult phenotype and reproductive success

The prenatal social environment affects offspring development in most studied taxa with potentially lifelong consequences. To understand the adaptive significance of such maternal influences on offspring development, it is important to study their effects on fitness. In guinea pigs, social instability during pregnancy leads to delayed development of male offspring. This has been interpreted as an adaptation to high social densities, where young males need to queue for reproductive opportunities since they cannot out-compete older dominant males. The consequences for male reproductive success are, however, so far unknown. To study the effects of different prenatal social densities on offspring reproductive performance, we housed females individually or in small groups during late pregnancy. Offspring from both treatments were reared together in large groups until independence and thereafter housed in same-sex pairs of the same treatment. We then observed courtship, aggressive behavior, and reproductive success in a low-density context with one male from each treatment competing over access to two females. Sons born to individually housed females initiated more fights, had more social contacts, courted females more, and had a higher reproductive success than sons of group-housed females. Sons born to mothers experiencing low social densities before birth therefore perform better at low social group sizes, suggesting that male development may be adaptively adjusted to anticipated social densities, although performance under high densities still needs to be compared.     

The number of males in primate social groups: a comparative test of the socioecological model

As applied to polygynous mammals, the socioecological model assumes that environmental risks and resources determine the spatial and temporal distribution of females, which then sets male strategies for monopolizing fertile matings. The effects of female spatial distribution (i.e., female number) and temporal overlap (female mating synchrony) have been examined in comparative studies of primates, but the relative influence of these two factors on male monopolization potential (the number of males) remains unclear. One particular problem is that female synchrony is more difficult to estimate than female number. This paper uses multivariate statistical methods and three independent estimates of female synchrony to assess the roles of spatial and temporal effects in the context of a phylogenetically corrected dataset. These analyses are based on sensitivity analyses involving a total of four phylogenies, with two sets of branch length estimates for each tree, and one nonphylogenetic analysis in which species values are used (because male behavior may represent a facultative response to the distribution of females). The results show: (1) that breeding seasonality predicts male number (statistically significant in six out of nine sensitivity tests); (2) that expected female overlap, after controlling for female group size using residuals, also accounts for the number of males in primate groups (significant in eight out of nine tests), and (3) that actual estimates of female mating synchrony predict male number, again after correcting for female group size (significant in five out of nine tests). Nonsignificant results are in the predicted direction, and female group size is significant in all statistical tests. These analyses therefore demonstrate an independent influence of female temporal overlap on male monopolization strategies in mammalian social systems. Received: 24 July 1998 / Received in revised form: 5 February 1999 / Accepted: 7 February 1999     

Reproductive success increases with degree of kinship in cooperative coalitions of female red howler monkeys (Alouatta seniculus)

Evaluation of evolutionary mechanisms proposed to promote cooperative behavior depends on the relative influence of the behavior on the reproductive success of individuals, the reproductive success of the group in which they interact behaviorally, and the degree of gene correlation among cooperators. The genetic relationship within cooperative coalitions of female red howler monkeys was examined for three populations with different densities and growth rates. Patterns of gene correlation change within coalitions is documented using data from the mitochondrial and nuclear genomes, and long-term census monitoring. Differences in fecundity and infant survivorship within and between groups of unrelated (rˉ=0) and related (rˉ≥ 0.25) females are compared. Females that emigrate from their natal groups form coalitions with other migrant females. These coalitions attempt to establish a territory and, once successful in producing offspring, exclude other females from feeding resources. Females in these coalitions had different mtDNA haplotypes and a genetically estimated mean r of 0, supporting demographic data on emigration patterns indicating that these females rarely have the opportunity to form coalitions with kin. Patterns of recruitment and rate of matriline development within social groups supported behavioral data indicating that females actively attempt to promote their own matriline as breeders over that of other females, and that some matrilines are more successful at this than others. Mean r among females was significantly higher in coalitions established as social groups for several generations (rˉ=0.44). In these groups, females all shared the same mtDNA haplotype, and mtDNA haplotype divergence was significantly higher between than within groups. Females in coalitions with kin had significantly higher reproductive success than females in unrelated coalitions in all populations. This difference was not a function of coalition size, number of males, socionomic sex ratio, or primiparity, although anecdotal evidence suggests that allomothering may compensate for inept new mothers in related coalitions more often than in unrelated ones. Differences in territory quality could not be ruled out as a potential causal factor in the saturated populations, but were unlikely in the low-density, growing population. There were substantial differences among long-established coalitions in overall reproductive output in all three populations, and this was significantly correlated with the number of breeding females. Increase in coalition size was a function of both group age and the behavioral tolerance among females. Regardless of the underlying reasons for the patterns observed, reproductive success clearly increases with degree of gene correlation among females within cooperative coalitions, and coalitions that recruit more daughters produce more offspring. The nature of the cooperative relationship among group females directly influences both of these outcomes. This is associated with substantial genetic differentiation among social groups within populations, creating conditions in which genetic tendencies towards cooperative behavior can become tightly associated with group reproductive success. Received: 15 September 1999 / Revised: 27 April 2000 / Accepted: 27 May 2000     

Cooperation in breeding by nonreproductive wrens: kinship,reciprocity, and demography

Summary Stripe-backed wrens (Campylorhynchus nuchalis) often live as adults in large groups on permanent, communally defended territories. Nonbreeding adults cooperate in rearing the young of a single breeding pair; this aid substantially increases the reproductive success of the breeders. In a 6-year study in Venezuela of a completely colorbanded population of 25–30 groups, most adults participated in breeding only as helpers and priority to breeding status was strictly age-determined. Detailed behavioral observations at breeding nests with nestlings showed that, in a sample of 100, helpers nearly always contributed as much to the care of young as breeders. Further, aid-giving does not vary systematically with relatedness of ycung to helpers or with probability of future reciprocation by young. Young being raised are most often at least half siblings of helpers, but seldom return aid to adults that helped raise them. Even adopted helpers collaborate fully. Patterns of demography and dispersal show slow turnover of breeders, delayed reproduction, and a viscous population structure.Application of Hamilton's condition for selection for aid-giving reveals that most individuals in this population can maximize inclusive fitness in the first 2 years by helping instead of breeding. Variation in helping effort and in age of first breeding is related to variation in natal group size and competition resulting from variable demographic neighborhoods in different years or in different parts of the population. Because reciprocation in the form of specific alliance formation among nonreproductives is uncommon, nonspecific reciprocity between cohorts and kin selection account well for the observed pattern of age-dependence in first breeding. Nondiscriminating helping in this population is associated with stable monogamous pair boncs, stable territory boundaries and group membership, strict seniortiy for breeding position, high viscosity and consistent effectiveness of aid. Under these circumstances, very simple behavioral rules amounting to nearly automatic helping seem sufficient to confer critical inclusive fitness gain on helpers.     

Brood parasitism and cooperative nesting in the moorhen,Gallinula chloropus

Summary During a two year study of the moorhen, Gallinula chloropus, a quarter of the nests found had two or more females laying in them. This was the result of two features of the moorhens' breeding biology: (1) Cooperative nesting; two (or more) females were paired to the same male. Both laid in the same nest, the second laid a large clutch synchronously to the first, and both cooperated in parental care. (2) Intraspecific brood parasitism; some females (parasites or dumpers) laid a small number of eggs at random in the nests' of neighbours. Cooperatively nesting females were mother and daughter, while parasite and host were not. Parasitic females laid in their own nest as well, and generally did so after completing a dumping sequence. Parasitic eggs were laid at random during the host's laying and incubation period, and were about a quarter as likely to produce an independent chick as were non-parasitic eggs. Parasitic females would probably have reared more chicks by laying in their own nests straight away, instead of dumping any eggs at all, because there was a strong decline in reproductive success with season. Possible reasons why they did not do so are discussed. Parasitised pairs tended to rear fewer own chicks than non-parasitised pairs. There was no evidence that hosts selectively destroyed parasitic eggs. Four hypotheses that explain why hosts accepted dumped eggs are considered. (1) The host pair were unaware of the dump, (2) the host male had copulated with the dumping female, (3) the dumper was related to one of the host pair, and (4) there was a benefit to the host pair because they gained extra helpers. These hypotheses were not supported by the data, but further tests are needed. As the cost of desertion was greater than the cost of acceptance of parasitic eggs, hosts may have been forced to accept.